Wolf

The wolf (Canis lupus[a]), also known as the gray wolf or grey
wolf, is a large canine native to Eurasia and North America. More Wolf
than thirty subspecies of Canis lupus have been recognized, and Temporal range:
gray wolves, as colloquially understood, comprise non- Middle Pleistocene–present
domestic/feral subspecies. The wolf is the largest extant member of (810,000–0 years BP) [1]
Canidae, males averaging 40 kg (88 lb) and females 37 kg (82 lb).
Wolves measure 105–160 cm (41–63 in) in length and 80–85 cm
(31–33 in) at shoulder height. The wolf is also distinguished from
other Canis species by its less pointed ears and muzzle, as well as a
shorter torso and a longer tail. The wolf is nonetheless related
closely enough to smaller Canis species, such as the coyote and the
golden jackal, to produce fertile hybrids with them. The banded fur
of a wolf is usually mottled white, brown, gray, and black, although
subspecies in the arctic region may be nearly all white.

Of all members of the genus Canis, the wolf is most specialized for
cooperative game hunting as demonstrated by its physical
adaptations to tackling large prey, its more social nature, and its Eurasian wolf (Canis lupus lupus) in
highly advanced expressive behaviour. It travels in nuclear families the Kolmården Wildlife Park, Sweden.
consisting of a mated pair accompanied by their offspring.
Offspring may leave to form their own packs on the onset of sexual 0:00 / 0:00
maturity and in response to competition for food within the pack.
Wolves are also territorial and fights over territory are among the
principal causes of wolf mortality. The wolf is mainly a carnivore Wolf pack howling
and feeds on large wild hooved mammals as well as smaller
Conservation status
animals, livestock, carrion, and garbage. Single wolves or mated
pairs typically have higher success rates in hunting than do large
packs. Pathogens and parasites, notably rabies, may infect wolves.

The global wild wolf population was estimated to be 300,000 in Least Concern (IUCN 3.1)[2]
2003 and is considered to be of Least Concern by the International
Union for Conservation of Nature (IUCN). Wolves have a long Scientific classification
history of interactions with humans, having been despised and Kingdom: Animalia
hunted in most pastoral communities because of their attacks on
livestock, while conversely being respected in some agrarian and Phylum: Chordata
hunter-gatherer societies. Although the fear of wolves exists in Class: Mammalia
many human societies, the majority of recorded attacks on people
have been attributed to animals suffering from rabies. Wolf attacks Order: Carnivora
on humans are rare because wolves are relatively few, live away
Family: Canidae
from people, and have developed a fear of humans because of their
experiences with hunters, ranchers, and shepherds. Genus: Canis
Species: C. lupus
Binomial name
Contents
Canis lupus
Etymology Linnaeus, 1758 [3]
Taxonomy Subspecies
Subspecies
Evolution See Subspecies of Canis lupus
Admixture with other canids
Description
Pelage
Ecology
Distribution and habitat Global wolf range based on the IUCN
Diet (2018).[2]
Interactions with other predators
Behaviour
Social structure
Reproduction
Hunting
Tracking
Pursuit
Disablement
Feeding
Infections
Viral and bacterial
Parasitic
Status and conservation
North America
Eurasia
Relationships with humans
In culture
In folklore, religion and mythology
In fable and literature
In heraldry and emblems
Conflicts
Wolf predation on livestock
Conflicts with dogs
Wolf attacks on humans
Human hunting of wolves
As pets and working animals
Notes
References
Bibliography
External links

Etymology
The English "wolf" stems from the Old English wulf, which is itself thought to be derived from the Proto-
Germanic *wulfaz. The Proto-Indo-European root *wĺ̥kʷos may also be the source of the Latin word for the
animal lupus (*lúkʷos).[4][5] The name "gray wolf" refers to the grayish colour of the species.[6]

Since pre-Christian times, Germanic peoples such as the Anglo-Saxons took on wulf as a prefix or suffix in
their names. Examples include Wulfhere ("Wolf Army", or "He whose army is the wolf"), Cynewulf ("Royal
Wolf"), Cēnwulf ("Bold Wolf"), Wulfheard ("Wolf-hard"), Earnwulf ("Eagle Wolf"), Wulfstān ("Wolf Stone")
Æðelwulf ("Noble Wolf"), Wolfhroc ("Wolf-Frock"), Wolfhetan ("Wolf Hide"), Isangrim ("Gray Mask"),
Scrutolf ("Garb Wolf"), Wolfgang ("Wolf Gait") and Wolfdregil ("Wolf Runner").[7]

Taxonomy
Canine phylogeny with ages of divergence

Gray wolf

1.10 mya
Coyote
1.32 mya

1.62 mya
African golden wolf
1.92 mya

2.74 mya

3.06 mya Ethiopian wolf

3.50 mya Golden jackal

Dhole

African wild dog

Side-striped jackal

2.62 mya
Black-backed jackal

Cladogram and divergence of the gray wolf (including the domestic dog) among its closest extant
relatives[8]
In 1758, the Swedish botanist and zoologist Carl Linnaeus published in his Systema Naturae the binomial
nomenclature.[3] Canis is the Latin word meaning "dog",[9] and under this genus he listed the doglike
carnivores including domestic dogs, wolves, and jackals. He classified the domestic dog as Canis familiaris,
and the wolf as Canis lupus.[3] Linnaeus considered the dog to be a separate species from the wolf because of
its cauda recurvata—its upturning tail—which is not found in any other canid.[10]

Subspecies

In the third edition of Mammal Species of the World published in 2005, the mammalogist W. Christopher
Wozencraft listed under C. lupus 36 wild subspecies, and proposed two additional subspecies: familiaris
(Linnaeus, 1758) and dingo (Meyer, 1793). Wozencraft included hallstromi—the New Guinea singing dog—
as a taxonomic synonym for the dingo. Wozencraft referred to a 1999 mitochondrial DNA study as one of the
guides in forming his decision, and listed the 38 subspecies of C. lupus under the biological common name of
"wolf", the nominate subspecies being the Eurasian wolf (C. l. lupus) based on the type specimen that
Linnaeus studied in Sweden.[11] Studies using paleogenomic techniques reveal that the modern wolf and the
dog are sister taxa, as modern wolves are not closely related to the population of wolves that was first
domesticated.[12] In 2019, a workshop hosted by the IUCN/Species Survival Commission's Canid Specialist
Group considered the New Guinea singing dog and the dingo to be feral dogs Canis familiaris, and therefore
should not be assessed for the IUCN Red List.[13]

Evolution

The phylogenetic descent of the extant wolf C. lupus from C. etruscus

through C. mosbachensis is widely accepted.[14] The earliest fossils
of C. lupus were found in what was once eastern Beringia at Old
Crow, Yukon, Canada, and at Cripple Creek Sump, Fairbanks,
Alaska. The age is not agreed upon but could date to one million
years ago. Considerable morphological diversity existed among
wolves by the Late Pleistocene. They had more robust skulls and
teeth than modern wolves, often with a shortened snout, a pronounced
development of the temporalis muscle, and robust premolars. It is
proposed that these features were specialized adaptations for the
processing of carcass and bone associated with the hunting and
scavenging of Pleistocene megafauna. Compared with modern
wolves, some Pleistocene wolves showed an increase in tooth
breakage similar to that seen in the extinct dire wolf. This suggests
they either often processed carcasses, or that they competed with other
carnivores and needed to consume their prey quickly. Compared with
A Canis etruscus skull in the
those found in the modern spotted hyena, the frequency and location
Montevarchi Paleontological
of tooth fractures in these wolves indicates they were habitual bone
Museum
crackers.[15] In June 2019, the severed yet preserved head of a
Pleistocene wolf, dated to over 40,000 years ago, was found close to
the Tirekhtyakh River in Yakutia, Russia, near the Arctic Circle. The
head was about 16 in (41 cm) long, much bigger than a modern wolf's head.[16][17][18]

Genomic studies suggest modern wolves and dogs descend from a common ancestral wolf
population[19][20][21] that existed 20,000 years ago.[19] Studies in 2017 and 2018 found that the Himalayan
wolf is part of a lineage that is basal to other wolves and split from them 691,000–740,000 years ago.[22][23]
Other wolves appear to have originated in Beringia in an expansion that was driven by the huge ecological
changes during the close of the Late Pleistocene.[23] A study in 2016 indicates that a population bottleneck
was followed by a rapid radiation from an ancestral population at a time during, or just after, the Last Glacial
Maximum. This implies that the original morphologically diverse wolf populations were out-competed and
replaced by more modern wolves.[24]
A 2016 genomic study suggests that Old World and New World wolves split around 12,500 years ago
followed by the divergence of the lineage that led to dogs from other Old World wolves around 11,100–
12,300 years ago.[21] An extinct Late Pleistocene wolf may have been the ancestor of the dog,[25][15] with the
dog's similarity to the extant wolf being the result of genetic admixture between the two.[15] The dingo,
Basenji, Tibetan Mastiff and Chinese indigenous breeds are basal members of the domestic dog clade. The
divergence time for wolves in Europe, the Middle East, and Asia is estimated to be fairly recent at around
1,600 years ago. Among New World wolves, the Mexican wolf diverged around 5,400 years ago.[21]

Admixture with other canids

In the distant past, there has been gene flow between

African golden wolves, golden jackals, and gray
wolves. The African golden wolf is a descendant of
a genetically admixed canid of 72% wolf and 28%
Ethiopian wolf ancestry. One African golden wolf
from the Egyptian Sinai Peninsula shows admixture
with Middle Eastern wolves and dogs.[26] There is Wolf–dog hybrids in the wild animal park at Kadzidłowo,
evidence of gene flow between golden jackals and Poland. Left: product of a male wolf and a female
Middle Eastern wolves, less so with European and spaniel; right: from a female wolf and a male West
Asian wolves, and least with North American Siberian Laika.
wolves. This indicates the golden jackal ancestry
found in North American wolves may have
occurred before the divergence of the Eurasian and North American wolves.[27]

The common ancestor of the coyote and the wolf has admixed with a ghost population of an extinct
unidentified canid. This canid is genetically close to the dhole and evolved after the divergence of the African
hunting dog from the other canid species. The basal position of the coyote compared to the wolf is proposed to
be due to the coyote retaining more of the mitochondrial genome of this unidentified canid.[26] Similarly, a
museum specimen of a wolf from southern China collected in 1963 showed a genome that was 12–14%
admixed from this unknown canid.[28] In North America, most coyotes and wolves show varying degrees of
past genetic admixture. The red wolf of the southeastern United States is a hybrid animal with 40%:60% wolf
to coyote ancestry. In addition, there was found to be 60%:40% wolf to coyote genetics in Eastern timber
wolves, and 75%:25% in the Great Lakes region wolves.[27]

In more recent times, some male Italian wolves originated from dog ancestry, which indicates female wolves
will breed with male dogs in the wild.[29] In the Caucasus Mountains, 10% of dogs including livestock
guardian dogs, are first generation hybrids.[30] Although mating between golden jackals and wolves has never
been observed, evidence of jackal-wolf hybridization was discovered through mitochondrial DNA analysis of
jackals living in the Caucasus Mountains[30] and in Bulgaria.[31]

Description
The wolf is the largest member of the Canidae family,[32] and is further distinguished from coyotes and jackals
by a broader snout, shorter ears, a shorter torso and a longer tail.[33][32] It is slender and powerfully built with
a large, deeply descending rib cage, a sloping back, and a heavily muscled neck.[34] The wolf's legs are
moderately longer than those of other canids, which enables the animal to move swiftly, and to overcome the
deep snow that covers most of its geographical range in winter.[35] The ears are relatively small and
triangular.[34] The wolf's head is large and heavy, with a wide forehead, strong jaws and a long, blunt
muzzle.[36] The skull is 230–280 mm (9–11 in) in length and 130–150 mm (5–6 in) in width.[37] The teeth are
heavy and large, making them better suited to crushing bone than those of other canids. They are not as
specialized as those found in hyenas though.[38][39] Its molars have a
flat chewing surface, but not to the same extent as the coyote, whose
diet contains more vegetable matter.[40] Females tend to have
narrower muzzles and foreheads, thinner necks, slightly shorter legs,
and less massive shoulders than males.[41]

Adult wolves measure 105–

160 cm (41–63 in) in length
and 80–85 cm (31–33 in) at
shoulder height.[36] The tail
measures 29–50 cm (11–
20 in) in length, the ears 90–
110 mm (31 ⁄2 –43 ⁄8 in) in
height, and the hind feet are
220–250 mm
5 7
(8 ⁄8 –9 ⁄8 in). [42] The size A North American wolf
A wolf skeleton housed in the Wolf
Museum, Abruzzo National Park, and weight of the modern
Italy wolf increases proportionally
with latitude in accord with Bergmann's rule.[43] The mean body
mass of the wolf is 40 kg (88 lb), the smallest specimen recorded at
12 kg (26 lb) and the largest at 79.4 kg (175 lb).[44][36] On average, European wolves weigh 38.5 kg (85 lb),
North American wolves 36 kg (79 lb), and Indian and Arabian wolves 25 kg (55 lb).[45] Females in any given
wolf population typically weigh 2.3–4.5 kg (5–10 lb) less than males. Wolves weighing over 54 kg (119 lb)
are uncommon, though exceptionally large individuals have been recorded in Alaska and Canada.[46] In
middle Russia, exceptionally large males are given a maximum weight of 69–79 kg (152–174 lb).[42]

Pelage

The wolf has very dense and fluffy winter fur, with a short undercoat
and long, coarse guard hairs.[36] Most of the undercoat and some
guard hairs are shed in spring and grow back in autumn.[45] The
longest hairs occur on the back, particularly on the front quarters and
neck. Especially long hairs grow on the shoulders and almost form a
crest on the upper part of the neck. The hairs on the cheeks are
elongated and form tufts. The ears are covered in short hairs and
project from the fur. Short, elastic and closely adjacent hairs are present
on the limbs from the elbows down to the calcaneal tendons.[36] The
Wolf in Spiti Valley, northern India
winter fur is highly resistant to the cold. Wolves in northern climates
can rest comfortably in open areas at −40 °C (−40 °F) by placing their
muzzles between the rear legs and covering their faces with their tail.
Wolf fur provides better insulation than dog fur and does not collect ice when warm breath is condensed
against it.[45]

In cold climates, the wolf can reduce the flow of blood near its skin to conserve body heat. The warmth of the
foot pads is regulated independently from the rest of the body and is maintained at just above tissue-freezing
point where the pads come in contact with ice and snow.[47] In warm climates, the fur is coarser and scarcer
than in northern wolves.[36] Female wolves tend to have smoother furred limbs than males and generally
develop the smoothest overall coats as they age. Older wolves generally have more white hairs on the tip of
the tail, along the nose, and on the forehead. Winter fur is retained longest by lactating females, although with
some hair loss around their teats.[41] Hair length on the middle of the back is 60–70 mm (23 ⁄8 –23 ⁄4 in), and the
guard hairs on the shoulders generally do not exceed 90 mm (31 ⁄2 in), but can reach 110–130 mm
(43 ⁄8 –51 ⁄8 in).[36]

A wolf's coat colour is determined by its guard hairs. Wolves usually

have some hairs that are white, brown, gray and black.[48] The coat of
the Eurasian wolf is a mixture of ochreous (yellow to orange) and
rusty ochreous (orange/red/brown) colours with light gray. The
muzzle is pale ochreous gray, and the area of the lips, cheeks, chin,
and throat is white. The top of the head, forehead, under and between
the eyes, and between the eyes and ears is gray with a reddish film.
The neck is ochreous. Long, black tips on the hairs along the back
form a broad stripe, with black hair tips on the shoulders, upper chest
and rear of the body. The sides of the body, tail, and outer limbs are a
pale dirty ochreous colour, while the inner sides of the limbs, belly,
Wolves in the La Boissière-du-Doré
and groin are white. Apart from those wolves which are pure white or
Zoo, France
black, these tones vary little across geographical areas, although the
patterns of these colours vary between individuals.[49]

In North America, the coat colours of wolves follow Gloger's rule, wolves in the Canadian arctic being white
and those in southern Canada, the U.S., and Mexico being predominantly gray. In some areas of the Rocky
Mountains of Alberta and British Columbia, the coat colour is predominantly black, some being blue-gray and
some with silver and black.[48] Differences in coat colour between sexes is absent in Eurasia;[50] females tend
to have redder tones in North America.[51] Black-coloured wolves in North America acquired their colour
from wolf-dog admixture after the first arrival of dogs that accompanied humans across the Bering Strait
12,000 to 14,000 years ago.[52] Research into the inheritance of white colour from dogs into wolves has yet to
be undertaken.[53]

Ecology

Distribution and habitat

Wolves occurred originally across Eurasia and North America.

Deliberate human persecution because of livestock predation and fear
of attacks on humans has reduced the wolf's range to about one-third
of what it once was. The wolf is now extirpated (locally extinct) in
much of Western Europe, the United States and Mexico, and in Japan.
In modern times, the wolf occurs mostly in wilderness and remote
areas. The wolf can be found between sea level and 3,000 m
(9,800 ft). Wolves live in forests, inland wetlands, shrublands, An Italian wolf in a mountainous
grasslands (including Arctic tundra), pastures, deserts, and rocky habitat in the Apennines in
peaks on mountains.[2] Habitat use by wolves depends on the Sassoferrato, Italy
abundance of prey, snow conditions, livestock densities, road
densities, human presence and topography.[40]

Diet
 Like all land mammals that are pack hunters, the wolf feeds predominantly on
wild herbivorous hoofed mammals that can be divided into large size 240–
650 kg (530–1,430 lb) and medium size 23–130 kg (51–287 lb), and have a
body mass similar to that of the combined mass of the pack members.[54][55]
The wolf specializes in preying on the vulnerable individuals of large prey,[40]
with a pack of 15 able to bring down an adult moose.[56] The variation in diet
between wolves living on different continents is based on the variety of
hoofed mammals and of available smaller and domesticated prey.[57]

In North America, the wolf's diet is dominated by wild large hoofed mammals
(ungulates) and medium-sized mammals. In Asia and Europe, their diet is
dominated by wild medium-sized hoofed mammals and domestic species. The
A wolf carrying a caribou wolf depends on wild species, and if these are not readily available, as in
hindquarter, Denali National Asia, the wolf is more reliant on domestic species.[57] Across Eurasia, wolves
Park, Alaska prey mostly on moose, red deer, roe deer and wild boar.[58] In North
America, important range-wide prey are elk, moose, caribou, white-tailed deer
and mule deer.[59] Wolves can digest their meal in a few hours and can feed
several times in one day, making quick use of large quantities of meat.[60] A well-fed wolf stores fat under the
skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.[61]

Nonetheless, wolves are not fussy eaters. Smaller-sized animals that may supplement their diet include rodents,
hares, insectivores and smaller carnivores. They frequently eat waterfowl and their eggs. When such foods are
insufficient, they prey on lizards, snakes, frogs, and large insects when available.[62] Wolves in northern
Minnesota prey on northern pike in freshwater streams.[63] The diet of coastal wolves in Alaska includes 20%
salmon,[64] while those of coastal wolves in British Columbia includes 25% marine sources, and those on the
nearby islands 75%.[65]

In Europe, wolves eat apples, pears, figs, melons, berries and cherries. In North America, wolves eat
blueberries and raspberries. Wolves also eat grass, which may provide some vitamins.[66] They are known to
eat the berries of mountain-ash, lily of the valley, bilberries, cowberries, European black nightshade, grain
crops, and the shoots of reeds.[62]

In times of scarcity, wolves will readily eat carrion.[62] In Eurasian areas with dense human activity, many
wolf populations are forced to subsist largely on livestock and garbage.[58] Prey in North America continue to
occupy suitable habitats with low human density, the wolves eating livestock and garbage only in dire
circumstances.[67] Cannibalism is not uncommon in wolves during harsh winters, when packs often attack
weak or injured wolves and may eat the bodies of dead pack members.[62][68][69]

Interactions with other predators

Wolves typically dominate other canid species in areas where they both occur. In North America, incidents of
wolves killing coyotes are common, particularly in winter, when coyotes feed on wolf kills. Wolves may attack
coyote den sites, digging out and killing their pups, though rarely eating them. There are no records of coyotes
killing wolves, though coyotes may chase wolves if they outnumber them.[70] According to a press release by
the U.S. Department of Agriculture in 1921, the infamous Custer Wolf relied on coyotes to accompany him
and warn him of danger. Though they fed from his kills, he never allowed them to approach him.[71]
Interactions have been observed in Eurasia between wolves and golden jackals, the latter's numbers being
comparatively small in areas with high wolf densities.[36][70][72] Wolves also kill red, Arctic and corsac foxes,
usually in disputes over carcasses, sometimes eating them.[36][73]
Brown bears typically dominate wolf packs in
disputes over carcasses, while wolf packs mostly
prevail against bears when defending their den
sites. Both species kill each other's young.
Wolves eat the brown bears they kill, while
brown bears seem to eat only young wolves.[74]
Wolf interactions with American black bears are
much rarer because of differences in habitat
preferences. Wolves have been recorded on
numerous occasions actively seeking out A wolf, a bear, coyotes and ravens compete over a kill in
American black bears in their dens and killing Yellowstone National Park.
them without eating them. Unlike brown bears,
American black bears frequently lose against
wolves in disputes over kills.[75] Wolves also dominate and sometimes kill wolverines, and will chase off those
that attempt to scavenge from their kills. Wolverines escape from wolves in caves or up trees.[76]

Wolves may interact and compete with felids, such as the Eurasian lynx, which may feed on smaller prey
where wolves are present[77] and may be suppressed by large wolf populations.[78] Wolves encounter cougars
along portions of the Rocky Mountains and adjacent mountain ranges. Wolves and cougars typically avoid
encountering each other by hunting at different elevations for different prey (niche partitioning). In winter,
when snow accumulation forces their prey into valleys, interactions between the two species become more
likely. Wolves in packs usually dominate cougars and can steal their kills or even kill them,[79] while one-to-
one encounters tend to be dominated by the cat. There are several documented cases of cougars killing
wolves.[80] Wolves more broadly affect cougar population dynamics and distribution by dominating territory
and prey opportunities and disrupting the feline's behaviour.[81] Wolf and Siberian tiger interactions are well-
documented in the Russian Far East, where tigers significantly depress wolf numbers, sometimes to the point
of localized extinction. Only human depletion of tiger numbers appears to protect wolves from competitive
exclusion from them. With perhaps only four proven records of tigers killing wolves, these cases are rare;
attacks appear to be competitive rather than predatory in nature.[82][77]

In Israel, Central Asia and India wolves may encounter striped hyenas, usually in disputes over carcasses.
Striped hyenas feed extensively on wolf-killed carcasses in areas where the two species interact. One-to-one,
hyenas dominate wolves, and may prey on them,[83] but wolf packs can drive off single or outnumbered
hyenas.[84][85] There is at least one case in Israel of a hyena associating and cooperating with a wolf pack. It is
proposed that the hyena could benefit from the wolves' superior ability to hunt large, agile prey. The wolves
could benefit from the hyena's superior sense of smell, to locate and dig out tortoises, to crack open large
bones, and to tear open discarded food containers like tin cans.[86]

Behaviour

Social structure

The wolf is a social animal.[36] Its populations consist of packs and lone wolves, most lone wolves being
temporarily alone while they disperse from packs to form their own or join another one.[87] The wolf's basic
social unit is the nuclear family consisting of a mated pair accompanied by their offspring.[36] The average
pack size in North America is eight wolves and in Europe 5.5 wolves.[43] The average pack across Eurasia
consists of a family of eight wolves (two adults, juveniles, and yearlings),[36] or sometimes two or three such
families,[40] with examples of exceptionally large packs consisting of up to 42 wolves being known.[88]
Cortisol levels in wolves rise significantly when a pack member dies,
indicating the presence of stress.[89] During times of prey abundance
caused by calving or migration, different wolf packs may join together
temporarily.[36]

Offspring typically stay in the pack for 10–54 months before

dispersing.[90] Triggers for dispersal include the onset of sexual
maturity and competition within the pack for food.[91] The distance
travelled by dispersing wolves varies widely; some stay in the vicinity
of the parental group, while other individuals may travel great Indian wolves at the Mysore Zoo
distances of upwards of 206 km (128 mi), 390 km (240 mi), and
670 km (420 mi) from their natal (birth) packs.[92] A new pack is
usually founded by an unrelated dispersing male and female, travelling together in search of an area devoid of
other hostile packs.[93] Wolf packs rarely adopt other wolves into their fold and typically kill them. In the rare
cases where other wolves are adopted, the adoptee is almost invariably an immature animal of one to three
years old, and unlikely to compete for breeding rights with the mated pair. This usually occurs between the
months of February and May. Adoptee males may mate with an available pack female and then form their own
pack. In some cases, a lone wolf is adopted into a pack to replace a deceased breeder.[88]

Wolves are territorial and generally establish territories far larger than they require to survive assuring a steady
supply of prey. Territory size depends largely on the amount of prey available and the age of the pack's pups.
They tend to increase in size in areas with low prey populations,[94] or when the pups reach the age of six
months when they have the same nutritional needs as adults.[95] Wolf packs travel constantly in search of prey,
covering roughly 9% of their territory per day, on average 25 km/d (16 mi/d). The core of their territory is on
average 35 km2 (14 sq mi) where they spend 50% of their time.[94] Prey density tends to be much higher on
the territory's periphery. Except out of desperation, wolves tend to avoid hunting on the fringes of their range
to avoid fatal confrontations with neighbouring packs.[96] The smallest territory on record was held by a pack
of six wolves in northeastern Minnesota, which occupied an estimated 33 km2 (13 sq mi), while the largest
was held by an Alaskan pack of ten wolves encompassing 6,272 km2 (2,422 sq mi).[95] Wolf packs are
typically settled, and usually leave their accustomed ranges only during severe food shortages.[36]

Wolves advertise their territories to other packs through howling and

scent marking. Scent marking involves urine, feces, and anal gland
scents. This is more effective at advertising territory than howling and
is often used in combination with scratch marks. Wolves increase their
rate of scent marking when they encounter the marks of wolves from
other packs. Lone wolves will rarely mark, but newly bonded pairs
will scent mark the most.[40] These marks are generally left every
240 m (260 yd) throughout the territory on regular travelways and
junctions. Such markers can last for two to three weeks,[95] and are
typically placed near rocks, boulders, trees, or the skeletons of large
A wolf marks its territory. Kolmården
animals.[36] Territorial fights are among the principal causes of wolf
Wildlife Park, Sweden.
mortality, one study concluding that 14–65% of wolf deaths in
Minnesota and the Denali National Park and Preserve were due to
other wolves.[97]

Wolves communicate to anticipate what their pack mates or other wolves might do next.[98] This includes the
use of vocalization, body posture, scent, touch, and taste.[99] The phases of the moon have no effect on wolf
vocalization, and despite popular belief, wolves do not howl at the moon.[100] Wolves howl to assemble the
pack usually before and after hunts, to pass on an alarm particularly at a den site, to locate each other during a
storm, while crossing unfamiliar territory, and to communicate across great distances.[101] Wolf howls can
under certain conditions be heard over areas of up to 130 km2 (50 sq mi).[40] Other vocalizations include
growls, barks and whines. Wolves do not bark as loudly or continuously as dogs do in confrontations, rather
barking a few times and then retreating from a perceived danger.[102] Aggressive or self-assertive wolves are
characterized by their slow and deliberate movements, high body posture and raised hackles, while submissive
ones carry their bodies low, flatten their fur, and lower their ears and tail.[103] Raised leg urination is
considered to be one of the most important forms of scent communication in the wolf, making up 60–80% of
all scent marks observed.[104]

Reproduction

Wolves are monogamous, mated pairs usually remaining together for

Korean wolves mating in the Tama
Zoological Park, Japan
life. Should one of the pair die, another mate is found quickly.[105]
With wolves in the wild, inbreeding does not occur where
outbreeding is possible.[106] Wolves become mature at the age of two
years and sexually mature from the age of three years.[105] The age of
first breeding in wolves depends largely on environmental factors:
when food is plentiful, or when wolf populations are heavily
managed, wolves can rear pups at younger ages to better exploit
abundant resources. Females are capable of producing pups every
year, one litter annually being the average.[107] Oestrus and rut begin
in the second half of winter and lasts for two weeks.[105]

Dens are usually constructed for pups during the summer period.
When building dens, females make use of natural shelters like fissures
in rocks, cliffs overhanging riverbanks and holes thickly covered by
vegetation. Sometimes, the den is the appropriated burrow of smaller
animals such as foxes, badgers or marmots. An appropriated den is
often widened and partly remade. On rare occasions, female wolves
dig burrows themselves, which are usually small and short with one to
three openings. The den is usually constructed not more than 500 m
(550 yd) away from a water source. It typically faces southwards
where it can be better warmed by sunlight exposure, and the snow
Iberian wolf pups stimulating their
can thaw more quickly. Resting places, play areas for the pups, and mother to regurgitate some food
food remains are commonly found around wolf dens. The odor of
urine and rotting food emanating from the denning area often attracts
scavenging birds like magpies and ravens. Though they mostly avoid areas within human sight, wolves have
been known to nest near domiciles, paved roads and railways.[108] During pregnancy, female wolves remain
in a den located away from the peripheral zone of their territories, where violent encounters with other packs
are less likely to occur.[109]

The gestation period lasts 62–75 days with pups usually being born in the spring months or early summer in
very cold places such as on the tundra. Young females give birth to four to five young, and older females from
six to eight young and up to 14. Their mortality rate is 60–80%.[110] Newborn wolf pups look similar to
German Shepherd Dog pups.[111] They are born blind and deaf and are covered in short soft grayish-brown
fur. They weigh 300–500 g (101 ⁄2 –173 ⁄4 oz) at birth and begin to see after nine to 12 days. The milk canines
erupt after one month. Pups first leave the den after three weeks. At one-and-a-half months of age, they are
agile enough to flee from danger. Mother wolves do not leave the den for the first few weeks, relying on the
fathers to provide food for them and their young. Pups begin to eat solid food at the age of three to four weeks.
They have a fast growth rate during their first four months of life: during this period, a pup's weight can
increase nearly 30 times.[110][112] Wolf pups begin play-fighting at the age of three weeks, though unlike
young coyotes and foxes, their bites are gentle and controlled. Actual fights to establish hierarchy usually
occur at five to eight weeks of age. This is in contrast to young coyotes and foxes, which may begin fighting
even before the onset of play behaviour.[113] By autumn, the pups are mature enough to accompany the adults
on hunts for large prey.[109]

Hunting

Single wolves or mated pairs typically have higher success rates in hunting than do large packs; single wolves
have occasionally been observed to kill large prey such as moose, bison and muskoxen unaided.[114][115] This
contrasts with the commonly held belief that larger packs benefit from cooperative hunting to bring down large
game.[115] The size of a wolf hunting pack is related to the number of pups that survived the previous winter,
adult survival, and the rate of dispersing wolves leaving the pack. The optimal pack size for hunting elk is four
wolves, and for bison a large pack size is more successful.[116]

As well as their physical adaptations for hunting hoofed mammals, wolves possess certain behavioural,
cognitive, and psychological adaptations to assist with their hunting lifestyle. Wolves are excellent learners that
match or outperform domestic dogs. They can use gaze to focus attention on where other wolves are looking.
This is important because wolves do not use vocalization when hunting. In laboratory tests, they appear to
exhibit insight, foresight, understanding, and the ability to plan. To survive, wolves must be able to solve two
problems—finding a prey animal, then confronting it.[117]

Tracking

Wolves move around their territory when hunting, using the same
trails for extended periods. After snowfalls, wolves find their old trails
and continue using them. These follow the banks of rivers, the
shorelines of lakes, through ravines overgrown with shrubs, through
plantations, or roads and human paths.[118] Wolves are nocturnal
predators. During the winter, a pack will commence hunting in the
twilight of early evening and will hunt all night, traveling tens of An Indian wolf trotting at Blackbuck
kilometres. Sometimes hunting large prey occurs during the day. National Park
During the summer, wolves generally tend to hunt individually,
ambushing their prey and rarely giving pursuit.[119]

The wolf usually travels at a loping pace, placing one of its paws directly in front of the other. This gait can be
maintained for hours at a rate of 8–9 km/h (5.0–5.6 mph).[120] On bare paths, a wolf can quickly achieve
speeds of 50–60 km/h (31–37 mph). The wolf has a running gait of 55–70 km/h (34–43 mph), can leap 5 m
(16 ft) horizontally in a single bound, and can maintain rapid pursuit for at least 20 minutes.[92] A wolf's foot is
large and flexible, which allows it to tread on a wide variety of terrain. A wolf's legs are long compared to
their body size allowing them to travel up to 76 km (47 mi) in 12 hours. This adaptation allows wolves to
locate prey within hours, but it can take days to find prey that can be killed without great risk. Moose and deer
live singly in the summer. Caribou live in herds of thousands which presents dangers for wolves. Elk live in
small herds and these are a safer target.[117]

A wolf carries its head at the same level as its back, lifting it only when alert.[36] In one study, wolves detected
moose using scent ten times, vision six times, and once by following tracks in the snow. Their vision is as
good as a human's, and they can smell prey at least 2.4 km (11 ⁄2 mi) away. One wolf travelled to a herd
103 km (64 mi) away. A human can detect the smell of a forest fire over the same distance from downwind.
The wolf's sense of smell is at least comparable to that of the domestic dog, which is at least 10,000 times more
sensitive than a human's.[117]
Pursuit

When hunting large gregarious prey, wolves will try to isolate an

individual from its group.[121] If successful, a wolf pack can bring
down game that will feed it for days, but one error in judgement can
lead to serious injury or death. Most large prey have developed
defensive adaptations and behaviours. Wolves have been killed while
attempting to bring down bison, elk, moose, muskoxen, and even by
one of their smallest hoofed prey, the white-tailed deer. With smaller
prey like beaver, geese, and hares, there is no risk to the wolf.[117]
Although people often believe that wolves can easily overcome any
An American bison in Yellowstone of their prey, their success rate in hunting hoofed prey is usually
National Park. Bison, elk, and moose low.[122]
usually stand their ground against a
pack of wolves. Generally, bison, elk, and moose will stand their ground, then the
wolves must struggle with them to bring them down. Often caribou
and deer will flee, but sometimes deer also make a stand.[117] If the
targeted animal stands its ground, wolves either ignore it, or try to intimidate it into running.[114] Wolves, or
even a wolf on its own, will attempt to frighten a herd into panicking and dispersing.[123]

When wolves encounter prey that flees, they give chase. The speed of sprinting prey is closely related to the
speed of their main predators. Wolves can run at 56–64 km/h (35–40 mph) across several kilometres and will
often pursue prey for at least 1 km (1 ⁄2 mi). One wolf chased a caribou for 8 km (5 mi), another chased and
tracked a deer for 20 km (12 mi), and one 11-year-old wolf chased and caught an Arctic hare after seven
minutes. Most wolf prey will try to run to water, where they will either escape or be better placed to attempt to
ward off the wolves.[117]

Disablement

The wolf must give chase and gain on its fleeing prey, slow it down by biting through thick hair and hide, and
then disable it enough to begin feeding.[117] After chasing and then confronting a large prey animal, the wolf
makes use of its 6 cm (21 ⁄2 in) fangs and its powerful masseter muscles to deliver a bite force of 28 kg/cm2
(400 lbf/in2 ), which is capable of breaking open the skulls of many of its prey animals. The wolf leaps at its
quarry and tears at it. One wolf was observed being dragged for dozens of metres attached to the hind leg of a
moose; another was seen being dragged over a fallen log while attached to a bull elk's nose.[116]

The most common point of wolf attacks on moose is the upper hind legs.[124][125][126] Hind leg wounds are
inflicted from the rear, midway up the hock with the canine teeth. These leave gaping skin perforations over
4 cm (11 ⁄2 in) in diameter. Although blood loss, muscle damage, and tendon exposure may occur, there is no
evidence of hamstringing. Attacks also occur on the fleshy nose, the back and sides of the neck, the ears, and
the perineum.[126] Wolves may wound large prey and then lie around resting for hours before killing it when it
is weaker due to blood loss, thereby lessening the risk of injury to themselves.[127]

With medium-sized prey, such as roe deer or sheep, wolves kill by biting the throat, severing nerve tracks and
the carotid artery, thus causing the animal to die within a few seconds to a minute. With small, mouselike prey,
wolves leap in a high arc and immobilize it with their forepaws.[128] When prey is vulnerable and abundant,
wolves may occasionally surplus kill. Such instances are common with domestic animals, but rare with wild
prey. In the wild, surplus killing occurs primarily during late winter or spring, when snow is unusually deep
(thus impeding the movements of prey)[129] or during the denning period, when den bound wolves require a
ready supply of meat.[130] Medium-sized prey are especially vulnerable to surplus killing, as the swift throat-
biting method allows wolves to kill one animal quickly and move on to another.[128]
Feeding

Once prey is brought down, wolves begin to feed excitedly, ripping

and tugging at the carcass in all directions, and bolting down large
chunks of it.[131] The breeding pair typically monopolizes food to
continue producing pups. When food is scarce, this is done at the
expense of other family members, especially non-pups.[132] The
breeding pair typically eats first. They usually work the hardest at
killing prey, and may rest after a long hunt and allow the rest of the
family to eat undisturbed. Once the breeding pair has finished eating, Two wolves feeding on a white-tailed
the rest of the family tears off pieces of the carcass and transports deer
them to secluded areas where they can eat in peace. Wolves typically
commence feeding by consuming the larger internal organs, like the
heart, liver, lungs, and stomach lining. The kidneys and spleen are eaten once they are exposed, followed by
the muscles.[133] A wolf can eat 15–19% of its body weight in a single feeding.[61]

Infections

Viral and bacterial

Viral diseases carried by wolves include: rabies, canine distemper,

canine parvovirus, infectious canine hepatitis, papillomatosis, and
canine coronavirus.[134] Wolves are a major host for rabies in Russia,
Iran, Afghanistan, Iraq and India.[135] In wolves, the incubation
period is eight to 21 days, and results in the host becoming agitated,
deserting its pack, and travelling up to 80 km (50 mi) a day, thus
increasing the risk of infecting other wolves. Infected wolves do not Play media
show any fear of humans, most documented wolf attacks on people Footage of a wolf taken from
being attributed to rabid animals. Although canine distemper is lethal Abruzzo Natural Park showing
in dogs, it has not been recorded to kill wolves, except in Canada and advanced signs of canine distemper
Alaska. The canine parvovirus, which causes death by dehydration,
electrolyte imbalance, and endotoxic shock or sepsis, is largely
survivable in wolves, but can be lethal to pups. Wolves may catch infectious canine hepatitis from dogs,
though there are no records of wolves dying from it. Papillomatosis has been recorded only once in wolves,
and likely does not cause serious illness or death, though it may alter feeding behaviours. The canine
coronavirus has been recorded in Alaskan wolves, infections being most prevalent in winter months.[134]

Bacterial diseases carried by wolves include: brucellosis, Lyme disease, leptospirosis, tularemia, bovine
tuberculosis,[136] listeriosis and anthrax.[135] Wolves can catch Brucella suis from wild and domestic reindeer.
While adult wolves tend not to show any clinical signs, it can severely weaken the pups of infected females.
Although lyme disease can debilitate individual wolves, it does not appear to significantly affect wolf
populations. Leptospirosis can be contracted through contact with infected prey or urine, and can cause fever,
anorexia, vomiting, anemia, hematuria, icterus, and death. Wolves living near farms are more vulnerable to the
disease than those living in the wilderness, probably because of prolonged contact with infected domestic
animal waste. Wolves may catch tularemia from lagomorph prey, though its effect on wolves is unknown.
Although bovine tuberculosis is not considered a major threat to wolves, it has been recorded to have killed
two wolf pups in Canada.[136]

Parasitic
Wolves carry ectoparasites and endoparasites; those in the former
Soviet Union have been recorded to carry at least 50 species.[135]
Most of these parasites infect wolves without adverse effects, though
the effects may become more serious in sick or malnourished
specimens.[137] Parasitic infection in wolves is of particular concern
to people. Wolves can spread them to dogs, which in turn can carry
the parasites to humans. In areas where wolves inhabit pastoral areas,
the parasites can be spread to livestock.[135]

Wolves are often infested with a variety of arthropod exoparasites,

A wolf in the Yellowstone National
including fleas, ticks, lice, and mites. The most harmful to wolves,
Park afflicted with sarcoptic mange
particularly pups, is the mange mite (Sarcoptes scabiei),[137] though
they rarely develop full-blown mange, unlike foxes.[36] Lice, such as
Trichodectes canis, may cause sickness in wolves, but rarely death. Ticks of the genus Ixodes can infect
wolves with Lyme disease and Rocky Mountain spotted fever.[137] The tick Dermacentor pictus also infests
wolves. Other ectoparasites include chewing lice, sucking lice and the fleas Pulex irritans and
Ctenocephalides canis.[36]

Endoparasites known to infect wolves include: protozoans and helminths (flukes, tapeworms, roundworms
and thorny-headed worms). Of 30,000 protozoan species, only a few have been recorded to infect wolves:
Isospora, Toxoplasma, Sarcocystis, Babesia, and Giardia.[137] Some wolves carry Neospora caninum, which
can be spread to cattle and is correlated with bovine miscarriages.[138] Among flukes, the most common in
North American wolves is Alaria, which infects small rodents and amphibians that are eaten by wolves. Upon
reaching maturity, Alaria migrates to the wolf's intestine, but does little harm. Metorchis conjunctus, which
enters wolves through eating fish, infects the wolf's liver or gall bladder, causing liver disease, inflammation of
the pancreas, and emaciation. Most other fluke species reside in the wolf's intestine, though Paragonimus
westermani lives in the lungs. Tapeworms are commonly found in wolves, as their primary hosts are
ungulates, small mammals, and fish, which wolves feed upon. Tapeworms generally cause little harm in
wolves, though this depends on the number and size of the parasites, and the sensitivity of the host. Symptoms
often include constipation, toxic and allergic reactions, irritation of the intestinal mucosa, and malnutrition.
Infections by the tapeworm Echinococcus granulosus in ungulate populations tend to increase in areas with
high wolf densities, as wolves can shed Echinoccocus eggs in their feces onto grazing areas.[137]

Wolves can carry over 30 roundworm species, though most roundworm infections appear benign, depending
on the number of worms and the age of the host. Ancylostoma caninum attaches itself on the intestinal wall to
feed on the host's blood, and can cause hyperchromic anemia, emaciation, diarrhea, and possibly death.
Toxocara canis, a hookworm known to infect wolf pups in the uterus, can cause intestinal irritation, bloating,
vomiting, and diarrhea. Wolves may catch Dioctophyma renale from minks, which infects the kidneys, and
can grow to lengths of 100 cm (40 in). D. renale causes the complete destruction of the kidney's functional
tissue and can be fatal if both kidneys are infected. Wolves can tolerate low levels of Dirofilaria immitis for
many years without showing any ill effects, though high levels can kill wolves through cardiac enlargement
and congestive hepatopathy. Wolves probably become infected with Trichinella spiralis by eating infected
ungulates. Although T. spiralis is not known to produce clinical signs in wolves, it can cause emaciation,
salivation, and crippling muscle pains in dogs. Thorny-headed worms rarely infect wolves, though three
species have been identified in Russian wolves: Nicolla skrjabini, Macrocantorhynchus catulinus, and
Moniliformis moniliformis.[137]

Status and conservation

The global wild wolf population in 2003 was estimated at 300,000.[139] Wolf population declines have been
arrested since the 1970s. This has fostered recolonization and reintroduction in parts of its former range as a
result of legal protection, changes in land use, and rural human population shifts to cities. Competition with
humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat
fragmentation pose a continued threat to the wolf. Despite these threats, the IUCN classifies the wolf as Least
Concern on its Red List due to its relatively widespread range and stable population. The species is listed by
the Convention on International Trade in Endangered Species of Wild Fauna and Flora in its Appendix II,
indicating that it is not threatened with extinction. However, those wolf populations living in Bhutan, India,
Nepal and Pakistan are listed in its Appendix I, indicating that these may become extinct without restrictions
on their trade.[2]

North America

In Canada, 50,000–60,000 wolves live in 80% of their historical

range, making Canada an important stronghold for the species.[40]
Under Canadian law, First Nations people can hunt wolves without
restrictions, but others must acquire licenses for the hunting and
trapping seasons. As many as 4,000 wolves may be harvested in
Canada each year.[140] The wolf is a protected species in national
parks under the Canada National Parks Act.[141] In Alaska, 7,000–
11,000 wolves are found on 85% of the state's 1,517,733 km2
Captive Mexican wolf at Sevilleta
(586,000 sq mi). Wolves may be hunted or trapped with a license; National Wildlife Refuge in New
around 1,200 wolves are harvested annually.[142] Mexico, as part of reintroduction

In the contiguous United States, wolf declines were caused by the

expansion of agriculture, the decimation of the wolf's main prey species like the American bison, and
extermination campaigns.[40] Wolves were given protection under the Endangered Species Act (ESA) of
1973, and have since returned to parts of their former range thanks to both natural recolonizations and
reintroductions.[143] Wolf populations in the Great Lakes states of Minnesota, Wisconsin and Michigan
number over 4,000 as of 2018.[144] Wolves also occupy much of the northern Rocky Mountains region, with
at least 1,704 wolves in Montana, Idaho and Wyoming as of 2015. They have also established populations in
Washington and Oregon.[145] In Mexico and parts of the southwestern United States, the Mexican and U.S.
governments collaborated from 1977 to 1980 in capturing all Mexican wolves remaining in the wild to prevent
their extinction and established captive breeding programs for reintroduction.[146] As of 2018, there were 230
Mexican wolves living in Mexico, 64 in Arizona, 67 in New Mexico, and 240 in captive breeding programs in
both countries.[147]

Eurasia

Europe, excluding Russia, Belarus and Ukraine, has 17,000 wolves in more than 28 countries.[149] In many
countries of the European Union, the wolf is strictly protected under the 1979 Berne Convention on the
Conservation of European Wildlife and Natural Habitats (Appendix II) and the 1992 Council Directive
92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora (Annex II and IV). There is
extensive legal protection in many European countries, although there are national exceptions.[2][150]
Protection and re-dispersal are strongly promoted by the Large Carnivore Initiative for Europe.

Wolves have been persecuted in Europe for centuries, having been exterminated in Great Britain by 1684, in
Ireland by 1770, in Central Europe by 1899, in France by the 1930s, and in much of Scandinavia by the early
1970s. They continued to survive in parts of Finland, Southern Europe, and Eastern Europe.[151] Since 1980,
European wolves have rebounded and expanded into parts of their former range. The decline of the traditional
pastoral and rural economies seems to have ended the need to
exterminate the wolf in parts of Europe.[140] As of 2016, estimates of
wolf numbers include: 4,000 in the Balkans, 3,460–3,849 in the
Carpathian Mountains, 1,700–2,240 in the Baltic states, 1,100–2,400
in the Italian peninsula, and around 2,500 in the northwest Iberian
peninsula as of 2007.[149]

In the former Soviet Union, wolf populations have retained much of

their historical range despite Soviet-era large scale extermination
campaigns. Their numbers range from 1,500 in Georgia, to 20,000 in
Kazakhstan and up to 45,000 in Russia.[152] In Russia, the wolf is
regarded as a pest because of its attacks on livestock, and wolf
management means controlling their numbers by destroying them
The wolf's range in Europe and
throughout the year. Russian history over the past century shows that
surrounding areas, as of 2019.
reduced hunting leads to an abundance of wolves.[153] The Russian
government has continued to pay bounties for wolves and annual
harvests of 20–30% do not appear to significantly affect their
numbers.[154]

During the 19th century,

wolves were widespread in
many parts of the Holy Land
east and west of the Jordan
River, but decreased
considerably in number
between 1964 and 1980,
largely due to persecution by
Definition of the conservation status
farmers.[155] In the Middle
of wolf populations according to the
East, only Israel and Oman
Large Carnivore Initiative for
give wolves explicit legal
Europe[148]
protection.[156] Israel has
A wolf in southern Israel protected its wolves since
1954 and has maintained a
moderately sized population of 150 through effective enforcement of
conservation policies. These wolves have moved into neighboring countries. Approximately 300–600 wolves
inhabit the Arabian Peninsula.[157] The wolf also appears to be widespread in Iran.[158] Turkey has an
estimated population of about 7,000 wolves.[159] Outside of Turkey, wolf populations in the Middle East may
total 1,000–2,000.[156]

In southern Asia, the northern regions of Afghanistan and Pakistan are important strongholds for wolves. The
wolf has been protected in India since 1972.[160] Hindus traditionally considered the hunting of wolves, even
dangerous ones, as taboo, for fear of causing a bad harvest. The Santals considered them fair game, as they did
every other forest-dwelling animal.[161] During British rule in India, wolves were not considered game
species, and were killed primarily in response to them attacking game herds, livestock, and people.[162] The
Indian wolf is distributed across the states of Gujarat, Rajasthan, Haryana, Uttar Pradesh, Madhya Pradesh,
Maharashtra, Karnataka and Andhra Pradesh.[163] As of 2019, it is estimated that there are around 2,000–
3,000 Indian wolves in the country.[164] In East Asia, Mongolia's population numbers 10,000–20,000. In
China, Heilongjiang has roughly 650 wolves, Xinjiang has 10,000 and Tibet has 2,000.[165] 2017 evidence
suggests that wolves range across all of mainland China.[166] Wolves have been historically persecuted in
China[167] but have been legally protected since 1998.[168] The last Japanese wolf was captured and killed in
1905.[169]
Relationships with humans

In culture

In folklore, religion and mythology

The wolf is a common motif in the mythologies and cosmologies of

peoples throughout its historical range. The Ancient Greeks
associated wolves with Apollo, the god of light and order.[170] The
Ancient Romans connected the wolf with their god of war and
agriculture Mars,[171] and believed their city's founders, Romulus and
Remus, were suckled by a she-wolf.[172] Norse mythology includes
the feared giant wolf Fenrir, eldest child of Loki and the giantess
Angrboda,[173] and Geri and Freki, Odin's faithful pets.[174] The wolf
The Capitoline Wolf, sculpture of the
was held in high esteem by the Dacians, who viewed it as the lord of
mythical she-wolf feeding the twins all animals and as the only effective defender against evil.[175]
Romulus and Remus, from the
legend of the founding of Rome, In the Pawnee creation myth, the wolf was the first animal brought to
Italy, 13th century AD. (The twins areEarth. When humans killed it, they were punished with death,
a 15th-century addition.) destruction and the loss of immortality.[176] For the Pawnee, Sirius is
the "wolf star" and its disappearance and reappearance signified the
wolf moving to and from the spirit world. Both the Pawnee and
Blackfoot call the Milky Way the "wolf trail".[177] The wolf is also an important totem animal for the Tlingit
and Tsimshian.[178] In Chinese astronomy, the wolf represents Sirius as the "blue beast" and the star itself is
called the "heavenly wolf".[179] In China, the wolf was traditionally associated with greed and cruelty and
wolf epithets were used to describe negative behaviours such as cruelty ("wolf's heart"), mistrust ("wolf's
look") and lechery ("wolf-sex").[180]

Tengrism places high importance on the wolf, as it is thought that, when

howling, it is praying to Tengri, thus making it the only creature other than
man to worship a deity.[181] In both Hinduism and Buddhism, the wolf is
ridden by gods of protection. In Vedic Hinduism, the wolf is a symbol of the
night and the daytime quail must escape from its jaws.[180] In Hindu
mythology, Krishna, to convince the people of Vraja to migrate to
Vṛndāvana, creates hundreds of wolves from his hairs, which frighten the
inhabitants of Vraja into making the journey.[182] In Tantric Buddhism,
wolves are depicted as inhabitants of graveyards and destroyers of
corpses.[180] In Zoroastrianism, the wolf has been demonized as the creation
of Ahriman.[183]

The concept of people turning into wolves, and the inverse, has been present
in many cultures. One Greek myth tells of Lycaon of Arcadia being Tlingit helmet and collar
transformed into a wolf by Zeus as punishment for his evil deeds. [184] The representing a wolf
legend of the werewolf has been widespread in European folklore and
involves people willingly turning into wolves to attack and kill others.[185]
The Navajo have traditionally believed that witches would turn into wolves by donning wolf skins and would
kill people and raid graveyards.[186] The Dena'ina believed wolves were once men and viewed them as
brothers.[170] Similarly, in Turkic mythology wolves were believed to be the ancestors of their people.[187]
In fable and literature

Aesop featured wolves in several of his fables, playing on the concerns of

Ancient Greece's settled, sheep-herding world. His most famous is the fable
of "The Boy Who Cried Wolf", which is directed at those who knowingly
raise false alarms, and from which the idiomatic phrase "to cry wolf" is
derived. Some of his other fables concentrate on maintaining the trust between
shepherds and guard dogs in their vigilance against wolves, as well as
anxieties over the close relationship between wolves and dogs. Although
Aesop used wolves to warn, criticize and moralize about human behaviour,
his portrayals added to the wolf's image as a deceitful and dangerous
animal.[188] The Bible contains 13 references to wolves, usually as metaphors
for greed and destructiveness. In the New Testament, Jesus is said to have
used wolves as illustrations of the dangers his followers, whom he represents Little Red Riding Hood
as sheep, would face should they follow him.(Matthew 7:15, Matthew 10:16 (1883), Gustave Doré
and Acts 20:29).[189] In the Jataka tales the wolf is portrayed as a trickster,
including one story in which it makes a hunter stop playing dead by pulling
on his club.[190] In another story, the wolf tricks and eats some rats by pretending to be lame but is foiled by
the chief rat.[191]

Isengrim the wolf, a character first appearing in the 12th-century Latin poem
Ysengrimus, is a major character in the Reynard Cycle, where he stands for
the low nobility, whilst his adversary, Reynard the fox, represents the peasant
hero. Isengrim is forever the victim of Reynard's wit and cruelty, often dying
at the end of each story.[192] The tale of "Little Red Riding Hood", first
written in 1697 by Charles Perrault, is considered to have further contributed
to the wolf's negative reputation in the Western world. The Big Bad Wolf is
portrayed as a villain capable of imitating human speech and disguising itself
with human clothing. The character has been interpreted as an allegorical
sexual predator.[193] Villainous wolf characters also appear in The Three Little
Pigs and "The Wolf and the Seven Young Goats".[194] The hunting of
wolves, and their attacks on humans and livestock, feature prominently in
Russian literature, and are included in the works of Leo Tolstoy, Anton
Akela, the Lone Wolf, leader Chekhov, Nikolay Nekrasov, Ivan Bunin, Leonid Pavlovich Sabaneyev, and
of the Seeonee Wolf Pack others. Tolstoy's War and Peace and Chekhov's Peasants both feature scenes
which raises Mowgli in The in which wolves are hunted with hounds and Borzois.[195] The musical Peter
Jungle Book. From an 1895 and the Wolf involves a wolf being captured for eating a farm duck, but is
illustration by John spared and sent to a zoo.[196]
Lockwood Kipling.
Wolves are among the central characters of Rudyard Kipling's The Jungle
Book. His portrayal of wolves has been praised posthumously by wolf
biologists for his depiction of them: rather than being villainous or gluttonous, as was common in wolf
portrayals at the time of the book's publication, they are shown as living in amiable family groups and drawing
on the experience of infirm but experienced elder pack members.[197] Farley Mowat's largely fictional 1963
memoir Never Cry Wolf is widely considered to be the most popular book on wolves, having been adapted
into a Hollywood film and taught in several schools decades after its publication. Although credited with
having changed popular perceptions on wolves by portraying them as loving, cooperative and noble, it has
been criticized for its idealization of wolves and its factual inaccuracies.[198][199][200] Lü Jiamin's 2004 semi-
autobiographical novel Wolf Totem uses wolves as symbols of freedom and independence. He associates the
Mongolian nomads with wolves and compares the Han Chinese of the present day to sheep, claiming they
accept any leadership. As such, the novel has caused controversy with the Chinese Communist Party.[201][202]
In heraldry and emblems

The wolf is a frequent charge in English heraldry. It is illustrated as a

supporter on the shields of Lord Welby, Rendel, and Viscount
Wolseley, and can be found on the coat of arms of Lovett and the vast
majority of the Wilsons and Lows. The demi-wolf is a common crest,
appearing in the arms and crests of members of many families,
including that of the Wolfes, whose crest depicts a demi-wolf holding
a crown in its paws, in reference to the assistance the family gave to
Charles II during the Battle of Worcester. Wolf heads are common in
The Pawnee Nation of Oklahoma
Scottish heraldry, particularly in the coats of Clan Robertson and
tribal flag
Skene. The wolf is the most common animal in Spanish heraldry and
is often depicted as carrying a lamb in its mouth, or across its
back.[203]

The wolf is featured on the flags of the Confederated Tribes of the Colville Reservation, the Oneida Nation of
Wisconsin and the Pawnee.[204] The Chechen wolf has been a symbol of the Chechen Nation.[205] In modern
times, the wolf is widely used as an emblem for military and paramilitary groups. It is the unofficial symbol of
the spetsnaz, and serves as the logo of the Turkish Gray Wolves. During the Yugoslav Wars, several Serb
paramilitary units adopted the wolf as their symbol, including the White Wolves and the Wolves of
Vučjak.[206]

Conflicts

Human presence appears to stress wolves, as seen by increased cortisol levels in instances such as
snowmobiling near their territory.[207]

Wolf predation on livestock

Livestock depredation has been one of the primary reasons for

A 1905 postcard of the Hexham wolf,
an escaped wolf shot for killing
livestock in England
North America, where healthy populations of wild prey have been largely restored.[208]
hunting wolves and can pose a severe problem for wolf conservation.
As well as causing economic losses, the threat of wolf predation
causes great stress on livestock producers, and no foolproof solution
of preventing such attacks short of exterminating wolves has been
found.[208] Some nations help offset economic losses to wolves
through compensation programs or state insurance.[209] Domesticated
animals are easy prey for wolves, as they have been bred under
constant human protection, and are thus unable to defend themselves
very well.[210] Wolves typically resort to attacking livestock when
wild prey is depleted. In Eurasia, a large part of the diet of some wolf
populations consists of livestock, while such incidents are rare in

The majority of losses occur during the summer grazing period, untended livestock in remote pastures being
the most vulnerable to wolf predation.[211] The most frequently targeted livestock species are sheep (Europe),
domestic reindeer (northern Scandinavia), goats (India), horses (Mongolia), cattle and turkeys (North
America).[208] The number of animals killed in single attacks varies according to species: most attacks on
cattle and horses result in one death, while turkeys, sheep and domestic reindeer may be killed in surplus.[212]
Wolves mainly attack livestock when the animals are grazing, though they occasionally break into fenced
enclosures.[213]
Conflicts with dogs

A review of the studies on the competitive effects of dogs on sympatric carnivores did not mention any
research on competition between dogs and wolves.[214][215] Competition would favour the wolf, which is
known to kill dogs; however wolves usually live in pairs or in small packs in areas with high human
persecution, giving them a disadvantage facing large groups of dogs.[215][216]

Wolves kill dogs on occasion, and some wolf populations rely on dogs as an important food source. In Croatia,
wolves kill more dogs than sheep, and wolves in Russia appear to limit stray dog populations. Wolves may
display unusually bold behaviour when attacking dogs accompanied by people, sometimes ignoring nearby
humans. Wolf attacks on dogs may occur both in house yards and in forests. Wolf attacks on hunting dogs are
considered a major problem in Scandinavia and Wisconsin.[208][217] The most frequently killed hunting breeds
in Scandinavia are Harriers, older animals being most at risk, likely because they are less timid than younger
animals, and react to the presence of wolves differently. Large hunting dogs such as Swedish Elkhounds are
more likely to survive wolf attacks because of their better ability to defend themselves.[217]

Although the number of dogs killed each year by wolves is relatively low, it induces a fear of wolves' entering
villages and farmyards to prey on them. In many cultures, dogs are seen as family members, or at least
working team members, and losing one can lead to strong emotional responses such as demanding more liberal
hunting regulations.[215]

Dogs that are employed to guard sheep help to mitigate human–wolf conflicts, and are often proposed as one
of the non-lethal tools in the conservation of wolves.[215][218] Shepherd dogs are not particularly aggressive,
but they can disrupt potential wolf predation by displaying what is to the wolf ambiguous behaviours, such as
barking, social greeting, invitation to play or aggression. The historical use of shepherd dogs across Eurasia
has been effective against wolf predation,[215][219] especially when confining sheep in the presence of several
livestock guardian dogs.[215][220] Shepherd dogs are sometimes killed by wolves.[215]

Wolf attacks on humans

The fear of wolves has been pervasive in many societies, though

humans are not part of the wolf's natural prey.[221] How wolves react
to humans depends largely on their prior experience with people:
wolves lacking any negative experience of humans, or which are
food-conditioned, may show little fear of people.[222] Although
wolves may react aggressively when provoked, such attacks are
mostly limited to quick bites on extremities, and the attacks are not
pressed.[221]

Predatory attacks may be preceded by a long period of habituation, in

which wolves gradually lose their fear of humans. The victims are Country children surprised by a wolf
repeatedly bitten on the head and face, and are then dragged off and (1833) by François Grenier de Saint-
consumed unless the wolves are driven off. Such attacks typically Martin
occur only locally and do not stop until the wolves involved are
eliminated. Predatory attacks can occur at any time of the year, with a
peak in the June–August period, when the chances of people entering forested areas (for livestock grazing or
berry and mushroom picking) increase.[221] Cases of non-rabid wolf attacks in winter have been recorded in
Belarus, Kirov and Irkutsk oblasts, Karelia and Ukraine. Also, wolves with pups experience greater food
stresses during this period.[36] The majority of victims of predatory wolf attacks are children under the age of
18 and, in the rare cases where adults are killed, the victims are almost always women.[221] Indian wolves
have a history of preying on children, a phenomenon called "child-lifting". They may be taken primarily in the
summer period in the evening hours, and often within human settlements.[223]
Cases of rabid wolves are low when compared to other species, as wolves do not serve as primary reservoirs
of the disease, but can be infected by animals such as dogs, jackals and foxes. Incidents of rabies in wolves are
very rare in North America, though numerous in the eastern Mediterranean, the Middle East and Central Asia.
Wolves apparently develop the "furious" phase of rabies to a very high degree. This, coupled with their size
and strength, makes rabid wolves perhaps the most dangerous of rabid animals.[221] Bites from rabid wolves
are 15 times more dangerous than those of rabid dogs.[224] Rabid wolves usually act alone, travelling large
distances and often biting large numbers of people and domestic animals. Most rabid wolf attacks occur in the
spring and autumn periods. Unlike with predatory attacks, the victims of rabid wolves are not eaten, and the
attacks generally occur only on a single day. The victims are chosen at random, though most cases involve
adult men. During the fifty years up to 2002, there were eight fatal attacks in Europe and Russia, and more
than two hundred in southern Asia.[221]

Human hunting of wolves

Theodore Roosevelt said wolves are difficult to hunt because of their

elusiveness, sharp senses, high endurance, and ability to quickly
incapacitate and kill a dog.[225] Historic methods included killing of
spring-born litters in their dens, coursing with dogs (usually
combinations of sighthounds, Bloodhounds and Fox Terriers),
poisoning with strychnine, and trapping.[226][227]

A popular method of wolf hunting in Russia involves trapping a pack

within a small area by encircling it with fladry poles carrying a human
Carcasses of hunted wolves in
scent. This method relies heavily on the wolf's fear of human scents,
Volgograd Oblast, Russia
though it can lose its effectiveness when wolves become accustomed
to the odor. Some hunters can lure wolves by imitating their calls. In
Kazakhstan and Mongolia, wolves are traditionally hunted with
eagles and falcons, though this practice is declining, as experienced falconers are becoming few in number.
Shooting wolves from aircraft is highly effective, due to increased visibility and direct lines of fire.[227] Several
types of dog, including the Borzoi and Kyrgyz Tajgan, have been specifically bred for wolf hunting.[215]

As pets and working animals

Wolves and wolf-dog hybrids are sometimes kept as exotic pets. Although closely related to domestic dogs,
wolves do not show the same tractability as dogs in living alongside humans, being generally less responsive
to human commands and more likely to act aggressively. A person is more likely to be fatally mauled by a pet
wolf or wolf-dog hybrid than by a dog.[228]

Notes
a. The domesticated dog (C.l. familiaris) and the dingo/New Guinea singing dog (C.l. dingo) are
included in the phylogenetic but not colloquial definition of 'wolf', and thus not in the scope of
this article.

References
1. Tedford, Richard H.; Wang, Xiaoming; Taylor, Beryl E. (2009). "Phylogenetic Systematics of the
North American Fossil Caninae (Carnivora: Canidae)". Bulletin of the American Museum of
Natural History. 325: 1–218. doi:10.1206/574.1 (https://doi.org/10.1206%2F574.1).
hdl:2246/5999 (https://hdl.handle.net/2246%2F5999).